A new genus and species of golden coral (Anthozoa, Octocorallia, Chrysogorgiidae) from the Northwest Atlantic

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1 ZooKeys 668: 1 10 (2017) A new genus and species of golden coral (Anthozoa, Octocorallia, Chrysogorgiidae)... 1 doi: /zookeys RESEARCH ARTICLE A peer-reviewed open-access journal Launched to accelerate biodiversity research A new genus and species of golden coral (Anthozoa, Octocorallia, Chrysogorgiidae) from the Northwest Atlantic Stephen D. Cairns 1, Ralf T. S. Cordeiro 2 1 Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, PO Box 37012, MRC 163, Washington, DC , USA 2 Programa de Pós-Graduação em Biologia Animal, Universidade Federal de Pernambuco, Av. Prof. Moraes Rego, 1235, Cidade Universitária, Recife, PE, CEP: , Brazil Corresponding author: Stephen D. Cairns (cairnss@si.edu) Academic editor: B.W. Hoeksema Received 9 February 2017 Accepted 2 April 2017 Published 12 April Citation: Cairns SD, Cordeiro RTS (2017) A new genus and species of golden coral (Anthozoa, Octocorallia, Chrysogorgiidae) from the Northwest Atlantic. ZooKeys 668: Abstract A new genus and species of unbranched golden coral, Flagelligorgia gracilis, is described based on several specimens collected off the southeastern coast of the United States. The genus is provisionally included in the family Chrysogorgiidae, pending molecular confirmation. Flagelligorgia morphologically resembles other unbranched chrysogorgiids, such as Distichogorgia, Chalcogorgia, Helicogorgia and Radicipes, to which it is compared. The type species is illustrated and its distribution mapped. Keywords Flagelligorgia gracilis, unbranched octocoral, biserial polyp arrangement, Radicipes, southeastern USA Introduction Chrysogorgiids form a polyphyletic family of octocorals (Pante et al. 2012), which is distributed in all oceans throughout the world (Watling et al. 2011). Most of its genera are in need of revision, some of which require reallocation into new families (Pante et al. 2012). In the course of doing a revision of the genus Radicipes Stearns, Copyright Stephen D. Cairns, Ralf T. S. Cordeiro. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

2 2 Stephen D. Cairns & Ralf T. S. Cordeiro / ZooKeys 668: 1 10 (2017) 1883 (see Cordeiro et al. in press), an unusual species was encountered that was similar to species in that genus that were already known, but was consistently different from them in several characters. Although no recently collected specimens were available for molecular analysis, sufficient alcohol-preserved specimens were present at the NMNH to describe the new species and the new genus in which it is placed. Material and methods All specimens are preserved in ethanol (70%) and deposited at the National Museum of Natural History. Reference material of all species of Radicipes was also examined (Cordeiro et al. in press). The terminology used in the description follows that of Bayer et al. (1983). Preparation of polyps for SEM to reveal sclerites includes short digestion (four seconds) of superficial coenenchyme, rinsing several times in distilled water, and drying at room temperature. Abbreviations used in the text include: Alb USFWS Albatross, L:W length to width ratio of a sclerite, MCC Monophyletic Chrysogorgiidae Clade sensu Pante et al. (2012), NMNH National Museum of Natural History, Smithsonian Institution, USNM United States National Museum (now known as the NMNH, but acronym still used for catalog numbers). Taxonomy Subclass Octocorallia Haeckel, 1866 Order Alcyonacea Lamouroux, 1812 Suborder Calcaxonia Grasshoff, 1999 Family Chrysogorgiidae Verrill, 1883 Chrysogorgidae Verrill 1883: 21. Chrysogorgiidae: Versluys 1902: 2 4; Bayer 1956: F216; Bayer and Muzik 1976: (key to genera); Bayer 1979: (key to genera); Cairns 2001: (synonymy); Pante and France 2010: 600 (key to genera); Watling et al. 2011: (distribution maps of all genera); Pante et al. 2012: 1 12 (phylogenetic and bathymetric analyses). Type genus. Chrysogorgia Duchassaing & Michelotti, Diagnosis (after Cairns (2001) and Pante et al. (2012)). Calcaxonians having an unjointed, solid (non-spicular), concentrically layered scleroproteinous axis. The axial layers are usually smooth (not undulated) and thus not longitudinally grooved externally; the axis usually displays metallic or iridescent reflections. The colony may be branched or unbranched (flagelliform), arising from a root-like or discoidal, strongly

3 A new genus and species of golden coral (Anthozoa, Octocorallia, Chrysogorgiidae)... 3 calcified holdfast. Polyps are contractile but not retractile, arranged in rows (uniserial, biserial or multiserial), but never in whorls. Sclerites predominantly flat, smooth scales, in some species warty rods and spindles. Remarks. Fourteen genera have been assigned to the Chrysogorgiidae, but based on sequencing of three genes, Pante et al. (2012) have suggested that only six of these genera belong to Chrysogorgiidae sensu stricto, which they call the Monophyletic Chrysogorgiidae Clade, or MCC. They imply that the other genera may belong to as many as three other, as yet undescribed, families. The six genera of the MCC are keyed by Pante and France (2010) and their worldwide distributions plotted by Watling et al. (2011). They are further discussed by Pante et al. (2012) in the context of a phylogenetic analysis. Flagelligorgia is morphologically most similar to Radicipes, which is part of the MCC, but lacking material suitable for molecular analysis, the placement of Flagelligorgia in that family can only be a suggestion at this time. In fact, we did attempt to sequence four genes (COI, mtmuts, 28S and 18S) from specimens collected in 1964 with no results. Distribution. Worldwide, m depths (Pante and Watling 2011: 6). Flagelligorgia gen. n. Type species. Flagelligorgia gracilis, here designated. Gender: feminine. Diagnosis. Colonies unbranched (flagelliform), loosely coiled, attached by a holdfast; axis composed of undulating concentric layers (Figs 1C E, 2D). Polyps arranged biserially. Body wall and coenenchymal scales elongate, smooth scales. Polyps lack an operculum. Remarks. Until 1956 (Bayer 1956) the Chrysogorgiidae had been divided into three subfamilies, one being the Lepidogorgiinae Versluys, 1902, consisting of genera with an unbranched colony and lacking polyp opercula. However, the subfamily classification has been ignored for decades and is not supported by molecular evidence. Nonetheless, it is incumbent to compare Flagelligorgia to the four other unbranched chrysogorgiid genera. Helicogorgia Bayer, 1981, known only from the east coast of Africa at m depth range, is unbranched but has its polyps arranged in a multiserial fashion on one side of the stem, a naked track displayed on the opposite side. Furthermore, its body wall scales are irregular plates and its coenenchymal scales are spindles. For all of these reasons Helicogorgia is easily distinguished from Flagelligorgia. Based on sequencing of three genes, Pante et al. (2012) indicate that Helicogorgia might constitute a family on its own, as sister to the Chrysogorgiidae. Helicogorgia was revised by Williams (1992), recognizing four species in the African coast. Chalcogorgia Bayer, 1949, known only from its type locality off northwestern Cuba at 708 m depth, is also unbranched and has biserially arranged polyps, like Flagelligorgia, but differs in having eight triangular opercular scales on each polyp, and body wall scales shaped as irregularly shaped platelets. According to Pante et al. (2012:

4 4 Stephen D. Cairns & Ralf T. S. Cordeiro / ZooKeys 668: 1 10 (2017) 8), based on very limited data, Chalcogorgia was suggested to ally with Helicogorgia, and thus also outside of the MCC. Distichogorgia Bayer, 1979, known only from the Blake Plateau off Florida at 814 m depth, is unbranched and has biserially arranged polyps, but differs in having unique body wall scales consisting of two fans of longitudinally arranged, blade-like scales, one on each side of the polyp, interspersed with tiny pointed sclerites, similar to its coenenchymals. Pante et al. (2012) did not report sequencing data for this genus, but based on its biserially polyps, they suggested an affinity with Helicogorgia and Chalcogorgia in a family separate from the Chrysogorgiidae. Perhaps most similar to Flagelligorgia is the genus Radicipes Stearns, 1883, consisting of 9 10 species (see Cordeiro et al., in press), which occurs worldwide at depths of m. Radicipes is unbranched but differs from Flagelligorgia in having uniserially arranged polyps, and a body wall and coenenchymal sclerites that are rod-shaped to compressed rod-shaped. Furthermore, chrysogorgiids (MCC), including Radicipes, are characterized by having an axis formed by non-undulated (smooth) concentric layers (Bayer 1956), which is not the case for Flagelligorgia. Based on molecular evidence Pante et al. (2012) placed Radicipes firmly in the Chrysogorgiidae clade (MCC), as the sister genus to Chrysogorgia. Lacking molecular evidence for Flagelligorgia (all specimens are quite old), it is not possible at this time to determine if it is allied with Radicipes in the Chrysogorgiidae (based on similarity of body wall sclerites) or the other cluster of genera including Helicogorgia, Chalcogorgia and Distichogorgia (based on polyp arrangement), which may constitute an as yet undescribed family. Distribution. Off Southeastern United States from North Carolina to Florida, m depths. Etymology. The name is a combination of flagellum (Latin for small whip) and gorgia (a common octocoral suffix and once the name of the order Gorgonacea, the name derived from the mythical female monster, the Gorgon), in reference to the flagelliform nature of the colony. Flagelligorgia gracilis sp. n. Figs 1 4 Material examined (Types). Holotype: Alb-2666, 1 specimen now in two pieces, USNM Paratypes: Alb-2601, 34 39'15"N, 33 30'10"W, 196 m, 1 colony, USNM 16607, 18 October 1885; Alb-2602, 34 38'30"N, 75 33'30"W, 227 m, 1 colony, USNM 16821, 18 October 1885; Alb-2666, 30 47'30"N, 79 49'W, 494 m, 33 colonies, SEM stubs , USNM 14458, 5 May 1886; Alb-2667, 30 53'N, 79 42'30"W, 499 m, 40 colonies, USNM 14457, 5 May 1886; Gerda-179, 27 41'N, 79 11'W, m, 9 colonies, USNM 57315, 1 July 1963; Megalopa, 11.2 km SSE of Carysfort Reef, Florida Keys, 205 m, 9 July 1950, 1 colony, USNM 51956, 9 July

5 A new genus and species of golden coral (Anthozoa, Octocorallia, Chrysogorgiidae)... 5 Figure 1. Diagnostic characters of Flagelligorgia gracilis sp. n. A general view of the holotype (USNM 49503) attached to a rocky substrate B detail of the holotype showing the biserial arrangement of polyps C,E cross-section of the stem of the holotype showing its four rounded longitudinal cords D detail of undulating concentric layers of the stem in cross-section. Scale bars: A: 10 mm, B: 5 mm, C, E: 0.1 mm, D: ; Pillsbury-105, 31 00'N, 79 42'W, m, 2 colonies, USNM 57316, 27 July 1964; Pillsbury-197, 27 59'N, 79 20'W, m, 2 colonies, USNM 52913, 11 August Type locality '30"N, 79 49'W (continental slope off Georgia), 494 m depth. Description. The colony is unbranched (flagelliform) and quite delicate, with an ascending clockwise spiral growth form (Fig. 1A, 2C). One of the largest specimens (the holotype) is 19 cm in length and only 1.1 mm in basal stem diameter. Colonies are attached to hard objects such as the deep-water coral Lophelia pertusa (Linnaeus, 1758), rocks, or echinoid tests, having a thin basal encrustation (holdfast) up to 7 mm in diameter; there are no root-like holdfasts. The axis is longitudinally grooved (Fig. 2D), resulting in four rounded longitudinal cords (1C, E), reflecting the underly-

6 6 Stephen D. Cairns & Ralf T. S. Cordeiro / ZooKeys 668: 1 10 (2017) Figure 2. Detailed view of polyps and axis in Flagelligorgia gracilis sp. n. (USNM 14458) through Scanning Electron Microscopy. A stereo view of polyps B oral view of a polyp C stereo view of a polyp s abaxial side D stereo view of the axis. Scale bars: A, C: 0.2 mm, B: 0.04 mm, D: 0.01 mm. ing undulated layers of scleroprotein (Fig. 1D), most easily seen in stem cross section. The axis is golden, the coenenchymal tissue usually pale brown to a dull yellow, in ethanol. Polyps are lacking from the proximal mm of the stem, which is approximately 1/5 1/3 of the colony length, depending on its maturity (Fig. 1A). Polyps occur biserially, on opposite sides of the stem in alternating fashion (Figs 1B, 2A, C), and are relatively closely spaced such that polyps occur per cm length. The polyps are mm in length and are somewhat cigar-shaped (Fig. 2C), being slightly tapered distally; the greatest diameter ( mm) is usually at mid-length. The body wall is covered with longitudinally oriented, elongate (L:W = ), imbricating scales (Fig. 3B) that measure mm in length. They are rounded distally, have smooth lateral edges, and are quite thin (e.g., µm in thickness). Their outer and inner surfaces are smooth. Toward the end of the polyp are smaller scales

7 A new genus and species of golden coral (Anthozoa, Octocorallia, Chrysogorgiidae)... 7 Figure 3. Sclerites of Flagelligorgia gracilis sp. n. (USNM 49503, holotype). A rods from coenenchyme B elongate rods from body wall C highly granular scales from first centimeters of the stem D tentacular scales. Scale bars: A C: 0.05 mm, D: 0.02 mm. associated with the tentacles (Figs 2B, 3D), similar in shape to the body wall scales but only mm in length. Pinnular sclerites are virtually absent. Scales show concentric bands of interference colors in polarized light. There are no sclerites in the

8 8 Stephen D. Cairns & Ralf T. S. Cordeiro / ZooKeys 668: 1 10 (2017) Figure 4. Distribution map of Flagelligorgia gracilis sp. n. axial sheath of coenenchyme. The outer coenenchymal scales (Fig. 3A) are also longitudinally arranged on the stem, elongate (L:W = ), and imbricate. They have pointed tips and their lateral edges are slightly serrate, each serration up to 5 µm in height; they are also quite thin, and their faces are also smooth. Coenenchymal scales on the first few centimeters of the stem are highly granular (Fig. 3C). Remarks. As in other unbranched octocorals, such as species of Radicipes, some specimens (e.g., from USNM 14458, USNM and USNM 14457) host commensal ophiuroids. Other octocoral species found at the same stations at which Flagelligorgia were collected include: Plumarella aurea (Deichmann, 1936), Plumarella pourtalesii (Verrill, 1883), Plumarella dichotoma Cairns & Bayer, 2004, Swiftia casta (Verrill, 1883) and Callogorgia americana Cairns & Bayer, Distribution. Southeastern coast of United States from off Outer Banks, North Carolina to off Carysfort Reef (near Key Largo, Florida) (Fig. 4), m depths. Etymology. Named gracilis (Latin for slender, gracile), in allusion to the very slender aspect of the colony. Acknowledgments We are grateful to the reviewers Dr. Eric Pante and Dr. Scott France and to Dr. Bert Hoeksema (editor), for their valuable suggestions for the improvement of this manuscript. The second author thanks the Smithsonian for a Short Term Visitor grant, which facilitated the accomplishment of this study. RTSC was also supported by FACEPE (IBPG /13), Brazil.

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